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Spatial variability that occurs at large scales has long been used by ecologists as a tool to examine the controls over ecosystem structure and function. Correlations of control variables such as climatic factors and response variables such as vegetation and soil carbon storage across broad regions have played a crucial role in predicting the response of ecosystems to global climate change. Despite the importance of these large-scale space-for-time substitutions, there are substantial limitations. One of these limitations is that many of the possible control factors covary with one another, and only some of the important control factors actually exist in large-scale databases. Thus, the true proximal controls may be difficult to identify. A second limitation is that models of spatial variability may not be appropriately applied to temporal variability. In this paper, we utilize a new approach to determine the extent to which N availability may constrain aboveground primary productivity in the Central Grassland region of the U.S. The strong relationship between average annual primary production and average annual precipitation found in spatial patterns in ecosystems globally has often been interpreted as evidence of a fundamental water limitation. However, temporal variation in annual aboveground net primary production (ANPP) indicates that other factors constrain production. We generated a spatial and temporal database for annual aboveground net primary production and annual net N mineralization by linking a database of input variables (precipitation, temperature, and soils) with predictive models. We generated independent data sets of aboveground net primary production and net N mineralization by using regression models to predict aboveground net primary production, and the Century model to simulate net N mineralization. Our analyses indicate that net primary production and net N mineralization both increase with mean annual precipitation; thus, it is not possible to separate the extent to which ANPP is controlled by water or N availability. Nitrogen use efficiency (NUE) increased with increasing precipitation across the region. Aboveground net primary production decreased with increasing temperature across the region, while N mineralization increased slightly, leading to decreasing (NUE) with increasing temperature. At high precipitation levels, aboveground net primary production increased and N mineralization decreased slightly with increasing soil fineness. Nitrogen use efficiency generally increased with increasing pools of soil organic matter, likely because in grasslands, the proportion of recalcitrant organic matter increases with the total organic matter pools. A comparison of interannual variation in net N mineralization with average spatial variation indicated a high degree of inertia in the response of N availability to precipitation levels. Our simulation results as well as field results of Lauenroth and Sala (1992) raise important questions about the applicability of space-for-time substitutions when dealing with ecosystem function. The structure of the systems appears to provide important constraints on the temporal variability that are not evident in an analysis of spatial variability.

We studied the effects of experimentally induced variation in leaf litter chemistry on leaf litter decomposition and leaf litter nutrient release of Carex species from habitats that differ in nutrient availability. Carex diandra, C. rostrata, and C. lasiocarpa are dominant in less productive mesotrophic fens, whereas C. acutiformis is dominant in more productive eutrophic fens. For each species, three types of litter were used: litter collected in the field (FLD); and litter from experimental populations grown at low (LN: 3.3 g N·m-2·yr-1) and high N supply (HN: 20.0 g N·m-2·yr-1). For all the litter chemistry parameters studied there were highly significant interactions between species and litter type. This implies that, due to differential plant-mediated controls on leaf litter chemistry, it is not always possible to predict changes in litter chemistry in response to increased nutrient supply. Litter decay was determined in a long-term (3 yr) field experiment using litter bags. The litter types of each species were incubated at their native growing sites. Contrary to what is generally found, the leaf litter decomposition rate of the species growing at the nutrient-poorest site (C. diandra) was higher than that of the species growing at the nutrient-richest site (C. acutiformis). Only for C. diandra and C. lasiocarpa was the decomposition rate of the litter from the HN treatment higher than that of the field litter. Thus, increased nutrient supply does not necessarily lead to higher litter decomposition rates. Nutrient controls on litter decay changed with time: initial litter decay (≤ 3 mo) was strongly controlled (high r2 values) by all P-related litter quality parameters, whereas long-term litter decay (> 1 yr) was most strongly related with the phenolics: N ratio, the phenolics: P ratio, the lignin: N ratio, and the C:N ratio. Our data suggest that high levels of atmospheric N deposition, such as in the study area (The Netherlands), may lead to a relative shortage of P in the plant-derived substrates for bacteria and fungi. As a result, P-related litter chemistry parameters exert a strong influence on litter decay. Our data did not support the hypothesis that high-nutrient species increase nutrient cycling due to the production of easily decomposable litter with high rates of nutrient release. The leaf litter from Carex acutiformis, the species from the high-productivity fens, decomposed more slowly than that of the other species, immobilized more N and P and had a longer period of net N and P immobilization. However, this species has a higher litter production than the other species and thereby increases the rate of nutrient cycling. At the intraspecific level, increased nutrient supply led to lower amounts of immobilized N and P and faster N and P release from litter in most species, and thereby to a higher rate of nutrient cycling. This positive feedback between nutrient supply rate and the rate of nutrient cycling is reinforced by the increase in litter production in response to increased nutrient supply.

Experiments using planktonic organisms revealed that the balance of radiant energy and available nutrients regulated herbivore growth rates through their effects on abundance and chemical composition of primary producers. Both algae and herbivores were energy limited at low light/nutrient ratios, but both were nutrient limited at high light/nutrient ratios. Herbivore growth increased with increasing light intensity at low values of the light/nutrient ratio due to increases in algal biomass, but growth decreased with increasing light at a high light/nutrient ratio due to decreases in algal quality. Herbivore production therefore was maximal at intermediate levels of the light/nutrient ratio. The results contribute to an understanding of mass transfer mechanisms in ecosystems and illustrate the importance of integration of energy-based and material-based currencies in ecology.

We examined the changes in leaf phenolic chemistry and insect herbivory from saplings of two temperate deciduous species, Liriodendron tulipifera (tulip poplar) and Cornus florida (dogwood), planted in five microenvironments in Gilmer County, Georgia, USA. The experimental design permitted comparisons between saplings grown in an open field, under shade cloth within the field, on the edge between field and forest, in forest understory, and within canopy gaps established within the forest. Half of the trees in each microenvironment were fertilized. Leaves from each tree were sampled at the end of the growing season and 1989 and 1990 and analyzed for toughness, percent dry mass, total phenolics, hydrolyzable tannins, condensed tannins, and insect herbivory (percent leaf area damaged). The shade-tolerant dogwood saplings contained higher levels of total phenolics and hydrolyzable tannins than the shade-intolerant tulip poplar saplings. Dogwood generally had lower levels of herbivory. These results support earlier studies suggesting that slow-growing, shade-tolerant species tend to have higher levels of phenolics and experienced lower levels of herbivory than fast growing, shade-intolerant species. However, dogwood leaves contained lower levels of condensed tannins and were as tough as tulip poplar leaves. Sunlight availability had a significant positive influence on levels of phenolics in both species. Leaf phenolics generally increased with greater insolation from forest to field and when sunlight was greater within field for forest habitats. However, the levels of tannins in dogwood saplings only dropped significantly in the deep shade of the forest. The similar levels of dogwood phenolics in most microenvironments are indicative of the relatively high photosynthetic efficiency of this species in reduced light environments. Overall, these results are consistent with carbon/nutrient balance theory that predicts trade-offs in the allocation of photosynthate from defense to growth as light declines. Levels of insect herbivory and total phenolics were inversely related for dogwood. However, the relationship with tannins was less apparent. Herbivory on tulip poplar was unrelated to changes in phenolics, possibly reflecting the greater chemical diversity of that species. Fertilization increased the biomass of both species, but had no apparent influence on levels of leaf phenolics or insect hervivory. The lack of a fertilization effect was unexpected in light of previous suggestions that fertilization results in reduced phenolics and increased herbivory.

The amounts of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), total chlorophyll (Chl), and total leaf nitrogen were measured in fully expanded, young leaves of wheat (Triticum aestivum L.), rice (Oryza sativa L.), spinach (Spinacia oleracea L.), bean (Phaseolus vulgaris L.), and pea (Pisum sativum L.). In addition, the activities of whole-chain electron transport and carbonic anhydrase were measured. All plants were grown hydroponically at different nitrogen concentrations. Although a greater than proportional increase in Rubisco content relative to leaf nitrogen content and Chl was found with increasing nitrogen supply for rice, spinach, bean, and pea, the ratio of Rubisco to total leaf nitrogen or Chl in wheat was essentially independent of nitrogen treatment. In addition, the ratio of Rubisco to electron transport activities remained constant only in wheat. Nevertheless, gas-exchange analysis showed that the in vivo balance between the capacities of Rubisco and electron transport in wheat, rice, and spinach remained almost constant, irrespective of nitrogen treatment. The in vitro carbonic anhydrase activity in wheat was very low and strongly responsive to increasing nitrogen content. Such a response was not found for the other C3 plants examined, which had 10- to 30-fold higher carbonic anhydrase activity than wheat at any leaf-nitrogen content. These distinctive responses of carbonic anhydrase activity in wheat were discussed in relation to CO2-transfer resistance and the in vivo balance between the capacities of Rubisco and electron transport.

Flavonoids produced by legume roots are signal molecules acting both as chemoattractants and nod gene inducers for the symbiotic Rhizobium partner. Combined nitrogen inhibits the establishment of the symbiosis. To know whether nitrogen nutrition could act at the level of signal production, we have studied the expression of flavonoid biosynthetic genes as well as the production of flavonoids in the roots of plants grown under nitrogen-limiting or nonlimiting conditions. We show here that growth of the plant under nitrogen-limiting conditions results in the enhancement of expression of the flavonoid biosynthesis genes chalcone synthase and isoflavone reductase and in an increase of root flavonoid and isoflavonoid production as well as in the Rhizobium meliloti nod gene-inducing activity of the root extract. These results indicate that in alfalfa (Medicago sativa L.) roots, the production of flavonoids can be influenced by the nitrogen nutrition of the plant.

Tamarind seed xyloglucan was partially degraded with a purified endoglucanase (endoV) from Trichoderma viride. Analysis by high-performance anion-exchange chromatography showed that this digest was composed of fragments consisting of 1 to 10 repeating oligosaccharide units ([xg]1-[xg]10). To study the adsorption of xyloglucan fragments to cellulose in detail, this digest was fractionated on BioGel P-6. Fragments were separated satisfactorily up to 5 repeating oligosaccharide units ([xg]5). The galactose substitution of the fragments increased with increasing molecular weight. The BioGel P-6 pools, as well as polymeric xyloglucan ([xg]00), were tested for their ability to interact with Avicel crystalline cellulose. Quantitative binding to cellulose occurred for sequences consisting of (at least) 4 repeating units. The adsorption of [xg]4 to Avicel was very high relative to that of [xg]00. The dimensions of these fragments were such that they could also penetrate the smaller pores of cellulose. Apparently, the effective surface area for the polymers is much smaller. Adsorption isotherms of [xg]00 and [xg]4 showed a pattern that is typical for polydisperse systems. However, the mechanisms underlying these patterns were different. At high xyloglucan concentrations, this polydispersity resulted in preferential adsorption of the larger molecules in the case of [xg]00 and a more extensive colonization of the smaller pores of cellulose in the case of [xg]4. The pH influenced the interaction between xyloglucan (fragments) and cellulose to only a small extent.

Data on soil nutrient availability for humid tropical forests are often reported, but are rarely integrated in an ecologically meaningful way with other measures of nutrient cycling. In this paper, estimates of soil nutrient availability and the inverse of litterfall nutrient concentrations (as an index of plant nutrient use) were compared, using data from 36 sites throughout the humid tropics, to determine if relationships exist between commonly used indices of nutrient cycling for plants and soils. Measures of both extractable and total soil P were significantly and positively correlated with the ratio of litterfall mass/P, particularly for montane tropical forests. Extractable soil P was also significantly correlated with litterfall mass for lowland humid tropical forests, explaining 58% of the variability in litterfall mass. A weak, albeit significant correlation was found between exchangeable soil Ca and litterfall mass/Ca, even though soil extraction techniques vary greatly. No significant relationship was found for total soil N, the most commonly measured soil N pool, and the inverse of litterfall N concentrations. The results suggest that our indices of soil P are related to litterfall processes, but that other measures, particularly total soil N, may not be as relevant to nutrient cycling by the vegetation.

A mechanistic model is introduced for resource competition between plant species on the basis of the equations of De Wit (1960). The zero-isocline diagrams that can be drawn using this model produce qualitative predictions about the plant features that contribute to the success of plants in nutrient-poor and nutrient-rich environments, respectively. The relationship between the model presented and the Lotka-Volterra and the Tilman model is discussed. An example is presented where the presented theory applies to observations in the field and to the results of competition experiments.